After completing development, the late third instar larva voids its alimentary canal contents in preparation of forming a cocoon and moves to an undisturbed place to spin a silk-like cocoon in which it pupates (Lyons, 1915; Karandikar and Munshi, 1950; Joseph, 1981) (Fig. 1).
The principal factors triggering pupation are declining levels of juvenile insect hormone (Grant, 1996). The silkin excreted for the cocoon is produced by the salivary glands (Strenger, 1973). The resulting cocoon consists of soft and moist silk-like material (Dryden, 1993), measures about 4 by 2 mm (Soulsby, 1982), is loosely spun and whitish in color (Dryden, 1993) and is coated with dust and debris because of its stickiness (Soulsby, 1982) which aids in camouflaging it perfectly (Karandikar and Munshi, 1950; Dryden, 1989). Flea cocoons can be found in soil, on vegetation, in carpets, under furniture, and on animal bedding (Dryden, 1993).
Three stages within the pupal cocoon
Within the pupal cocoon three distinct stages are found in the order Siphonaptera (to which the cat and the dog flea belong): The first is the U-shaped larval prepupa, the second is a true exarate pupa and the third is the preemergent adult, which has completed its pupal-imaginal molt but remains within the cocoon for varying lengths of time (Silverman et al., 1981) (see Preemerged Adult).
The U-shaped larva begins pupal development about 18 hours after the completion of the cocoon (Dryden and Smith, 1994). If the larva is disturbed prior to this time in form of gentle shifting of the larval medium (e.g.) it might emerge and exit the cocoon. In trials by Dryden and Smith (1994) over 40% of those larvae did not spin a second cocoon, but developed as naked flea pupae. Metzger and Rust (1997) observed shorter times for adults to emerge from naked pupae compared with emergence time from cocoons.
The importance of this early emergence which can also be produced in carpeted homes by the vigorous agitation of the carpet by vacuums with rotary beater bars is so far unknown (Dryden and Smith, 1994). Of importance for a successful formation of the cocoon is the orientation against a perpendicular structure. Less than 3% manage to spin an enclosing cocoon in the absence of a vertical surface, but over 95% of the larvae nevertheless survive to adulthood, thus demonstrating that the cocoon is not essential for the development of adults (Dryden and Smith, 1994).
Nevertheless it offers some benefits, e.g. protection from ant predation (Silverman and Appel, 1984), somewhat impeding adult emergence and offering protection from non-host-produced stimuli and thus minimising non-host-induced emergence (Silverman and Rust, 1985). It does not provide a barrier to water loss (Silverman and Rust, 1985) and also does not seem to be a barrier to insecticides (Dryden and Reid, 1993), even though cocoons placed in a carpet appeared to be highly tolerant to a variety of insecticides (Rust and Reierson, 1989). The survival of developing pupae in insecticide-treated households is caused by the lack of penetration of the carpet canopy by the insecticide (Dryden and Rust, 1994).
Prepupae and pupae are fairly resistant life stages (Silverman et al., 1981), but it is unknown how much of that protection is provided by the cocoon (Dryden and Smith, 1994).
Favourable conditions for pupating
At 27+/-2°C, females pupate within about 32 hours and males pupate an average 12.1 hours later within about 44 hours (Dryden and Smith, 1994). The degree of development within the cocoon at 16°C and 27°C was similar regardless of humidity (in trials of Silverman et al., 1981). Under moderate conditions the pupa was the immature stage most resistant to desiccation; 80% survived to adulthood at 2% RH and 27°C (Silverman et al., 1981). Exposure to 35°C during pupal development is uniformly lethal (Silverman et al., 1981). Exposure to 3°C for five days and 8°C for 20 days is lethal as well (Silverman and Rust, 1983).
Females develop into adults about 1.6 days faster than males (Dryden and Smith, 1994). In trials of Metzger and Rust (1997) male prepupae and pupae developed more slowly than females at each temperature, requiring 14-20% more time to develop. At 15.5°C, the mean number of days for the female and male pupae to develop was 19.5 (female) and 23.5 (male), respectively (Metzger, 1995), or even nine days earlier (Metzger and Rust, 1997) and at 26.7°C, the mean number of days was 5.2 (female) and 6.5 (male), respectively (Metzger, 1995), or two to five days earlier (Metzger and Rust, 1997).
Not only females, but also adults that developed from light prepupae in general, remained in their cocoons for a shorter time than adults from more robust prepupae with two explanations proposed: either because of emergence mechanisms triggered when water and food reserves drop below a critical level or because of a weaker cocoon produced by incompletely nourished prepupa that does not impede adult emergence (Silverman and Rust, 1985). When pupae were maintained at 24.4°C and 78% RH, adult C. felis began to emerge eight days after the initiation of pupal development and by day 13, all fleas emerged (Dryden, 1988).
The pupa is suggested to be the stage most likely to survive extended periods in cool dry climates (Silverman et al. 1981).
Summary see Box 3.
BOX 3. Cat flea pupae
- Dryden MW: Evaluation of certain parameters in the bionomics of Ctenocephalides felis felis (Bouché 1835). 1988, MS Thesis, Purdue University, West Lafayette
- Dryden MW: Biology of the cat flea, Ctenocephalides felis felis. Comp Anim Pract. 1989, 19, 23-7
- Dryden MW: Biology of fleas of dogs and cats. Comp Cont Educ Pract Vet. 1993, 15, 569-79
- Dryden MW, Reid B: Investigations of cat flea pupation, cocoon formation and the impact of pupae on a flea control program. In: Knapp FW, Potter M, Labore D (eds.): Proc Int Symp Ectoparas Pets. 1993, Lexington University Ky Press, Lexington, pp 27-29
- Dryden MW, Rust MK: The cat flea: biology, ecology and control. Vet Parasitol. 1994, 52, 1-19
- Dryden MW, Smith V: Cat flea (Siphonaptera: Pulicidae) cocoon formation and development of naked flea pupae. J Med Entomol. 1994, 31, 272-7
- Grant D: Flea biology and control. Vet Pract. 1996, 28, 7-8
- Joseph SA: Studies on the bionomics of Ctenocephalides felis orientis (Jordan 1925). Cheiron. 1981, 10, 275-80
- Karandikar KR, Munshi DM: Life history and bionomics of the cat flea, Ctenocephalides felis felis (Bouché). J Bombay Nat His Soc. 1950, 49, 169-77
- Lyons H: Notes on the cat flea (Ctenocephalides felis (Bouché)). Psyche. 1915, 22, 124-32
- Metzger ME: Photoperiod and temperature effects on the development of Ctenocephalides felis (Bouché) and studies on its chemical control in turfgrass. 1995, MS Thesis, University of California, Riverside
- Metzger ME, Rust MK: Effect of temperature on cat flea (Siphonaptera: Pulicidae) development and overwintering. J Med Entomol. 1997, 34, 173-8
- Rust MK, Reierson DA: Activity of insecticides against the preemerged adult cat flea in the cocoon (Siphonaptera: Pulicidae). J Med Entomol. 1989, 26, 301-5
- Silverman J, Appel AG: The pupal cocoon of the cat flea, Ctenocephalides felis (Bouché) (Siphonaptera: Pulicidae): a barrier to ant predation. Proc Entomol Soc Wash. 1984, 86, 660-3
- Silverman J, Rust MK: Some abiotic factors affecting the survival of the cat flea Ctenocephalides felis (Siphonaptera: Pulicidae). Environ Entomol. 1983, 12, 490-5
- Silverman J, Rust MK: Extended longevity of the pre-emerged adult cat flea (Siphonaptera: Pulicidae) and factors stimulating emergence from the pupal cocoon. Ann Entomol Soc Am. 1985, 78, 763-8
- Silverman J, Rust MK, Reierson DA: Influence of temperature and humidity on survival and development on the cat flea, Ctenocephalides felis (Siphonaptera: Pulicidae). J Med Entomol. 1981, 18, 78-83
- Soulsby EJL (ed.): Helminths, arthropods and protozoa of domesticated animals. 7th edn., 1982, Lea & Febiger, Philadelphia
- Strenger A: [About the nutritional physiology of the larva of Ctenocephalides felis felis.] B Zool Jahrb Syst Bd. 1973, 100, 64-80 [in German]